Habitat selection of wintering passerines in salt marshes of the German Wadden Sea

The salt marshes of the Wadden Sea are important wintering areas for some species of granivorous passerines, which have declined considerably since the 1960s. We investigated the habitat choice of all wintering passerines in eight study areas in German salt marshes with special consideration of human impact on these habitats. Granivorous species that almost exclusively winter in salt marshes, Shorelark (Eremophila alpestris), Snow Bunting (Plectrophenax nivalis) and Twite (Carduelis flavirostris) were concentrated in the lower salt marsh vegetation and in the driftlines, while all other species preferred the high upper salt marsh communities, although Rock Pipits (Anthus petrosus littoralis) fed in muddy areas along ditches. Shorelarks switched habitat in conditions where seeds were scarce to feed instead on arthropods in upper salt marshes. Intensively sheep-grazed upper salt marshes resemble lower salt marshes in their vegetation and were therefore mainly visited by Shorelarks, Snow Buntings and Twites. In winter, the driftline is preferred by the two former species, while in autumn and spring more birds foraged in the salt marshes. Twites prefer to feed mainly on seeds of Salicornia. Areas with S. europaea are visited mainly in late autumn and early winter, while areas with S. stricta are used throughout the winter because of a steady supply of seeds. Several years after embankment, polders are hardly used any more by the lower salt marsh species as the habitat changes into freshwater marshes. Large embankment projects since the early 1960s have included salt marshes and intertidal flats, and the resultant loss of habitat is responsible for the decline of lower salt marsh species. For other passerine species the effects of reclamation are unknown. The effects of intensified grazing on the wintering populations of Shorelark, Snow Bunting and Twite are still unresolved. Although grazing supports lower salt marsh vegetation, the seed production per plant is much lower there and some important seed producers hardly occur. Since grazing was reduced and embankment projects have been stopped, the salt marsh areas (especially lower salt marshes) have increased and so have the wintering populations of Shorelark, Snow Bunting and Twite. For the other species, the consequences of habitat changes are unknown, although it is suggested that reduced grazing will support them. Reducing the human impact on salt marshes will, in the long run, probably lead to a natural salt marsh with much variety in elevation and in its corresponding vegetation and bird communities. Meanwhile, management by grazing might be required in parts of the salt marshes.     

The impact of grazing on plant communities,plant populations and soil conditions on salt marshes

Grazing an abandoned salt marsh causes retrogressive succession, since mid salt-marsh communities change into lower salt-marsh communities. Grazing and mowing are compared in detail. Both management practices enhance species diversity in an abandoned salt marsh. This can be attributed to the removal of litter. The finding that lower salt-marsh species appear more with grazing than with mowing or abandoning is not related to a higher soil salinity as compared to mowing or abandoning, but probably to locally baring of the soil by grazing animals. Only species of pioneer or unstable environments seem to have a persistent seed bank, for other species seed dispersal seems to be a limiting factor for their establishment.Nomenclature follows Heukels & van Ooststroom (1977) for species; Westhoff & den Held (1969) for syntaxa.Mrs R. Rusthoven analyzed the soil samples, Mr E. Leeuwinga drawed the figures, and Mrs J. O'Brien corrected the English text.     

Habitat switching by dark-bellied brent geese Branta b. bernicla (L.) in relation to food depletion

Seasonal changes in the distribution and feeding behaviour of dark-bellied brent geese Branta b. bernicla (L.) and the biomass of their food plants were studied in three successive winters on the Norfolk coast. The data was used, in conjunction with published information, to show how depletion, productivity and mortality of food plants drive the pattern of habitat switching in this species. It is then possible to explain the habitat shifts observed over the last 35 years and predict future changes. On arrival, geese fed first on algal beds and then on salt marsh, grass and arable fields before returning to feed entirely on the salt marsh in spring. The biomass of green algae, and subsequently the salt marsh vegetation, declined during the autumn and this could be attributed to depletion through goose grazing and natural mortality. As depletion occurred the geese fed more intensively, for a greater percentage of time and with an increasing pace rate, the net result, however, was a declining intake rate (as measured by defaecation rate). The algal biomass at which the geese switched from the algal beds to salt marsh was consistent between years, with heavy storm-induced loss of algae in one year resulting in an earlier switch. That the timing of habitat switches may be explained by depletion of food plants was further supported by historical data: the number of brent geese wintering at the site has increased dramatically over the last 30–35 years and the time of switching from algal beds to salt marsh and from salt marsh to salt marsh and fields has become progressively earlier, as expected from the increased depletion. The expected further increase in brent goose numbers will increase the rate of depletion of intertidal vegetation so that the switches between habitats will be more rapid and the geese will move inland earlier and remain inland longer. The expected increase in the brent goose population will thus result in a disproportionate increase in the levels of conflict between brent geese and agriculture.     

Long-term changes of salt marsh communities by cattle grazing

Over a period of 9 years a grazing experiment was carried out in the mainland salt marsh of the Leybucht (Niedersachsen) with three stocking rates, namely, 0.5 ha^-1, 1 ha^-1, and 2 cattle ha^-1. These were also compared with an abandoned area. The results are based on sampling of the invertebrates in 1980, 1981, 1982, and 1988, and of the vegetation in 1980 and 1988. The rate of sedimentation is highest in the Puccinellia maritima-zone and decreases with the increase of stocking rates. The Elymus pycnanthus vegetation type becomes dominant in the higher salt marsh in the abandoned site. The canopy height decreases with increasing stocking rate, whereas a gradient in the structure of the vegetation develops with the lowest stocking rate. The population densities, the species-richness and the community diversity of invertebrates increases after the cessation of grazing. The high rate of sedimentation in the abandoned site promotes the immigration of species from higher salt marsh levels and adjacent grasslands, and eventually halotopophilous species and communities may disappear. On the other hand grazing reduces numerous species living both in or on upper parts of the vegetation or being sensitive to trampling by cattle. The community structure shows that the salt marsh ecosystem changed from a food web dominated by plant feeding animals to a food web dominated by animals foraging on detritus. The salt marsh management has to be differentiated into both ungrazed and lightly grazed areas (each 50%) or an overall grazing in large areas with less than 0.5 cattle ha^-1.     

Interactive effects of time and vegetation on reproduction of redshanks (Tringa totanus) breeding in Wadden Sea salt marshes

As shown for various species, nesting waders are non-randomly distributed on wetlands and preferentially select riparian nest-sites adjacent to limnic or marine waterbodies. Studying the redshank Tringa totanus, we tested the hypotheses that, in a coastal wader species which conceals its clutch in vegetation, predation and hatching success are affected by vegetation zonation, and that breeding in lower salt marsh areas has negative consequences for reproduction. We further predicted effects of timing of breeding and breeding experience/age of adults potentially reflected by egg biometrics both on nest-site selection and reproduction. Effects of vegetation, space, time and individual quality on hatching success of redshanks were studied in the German part of the Wadden Sea. Dominant plant species, vertical vegetation structure and nest concealment varied significantly between nests. Variation in nest concealment was relatively low: about 90% of clutches were classified as being well concealed. This variation was explainable by vegetation structure but not by vegetation composition at the nest-site, distance to shoreline, and time of clutch initiation. Vertical vegetation structure varied by dominant plant species but not by distance to shoreline and time of clutch initiation. Hatching success of clutches was low (10.6%) due to high predation (daily predation rate: 7.4%). Hatching success and duration of clutch survival were negatively and predation positively related to the date of clutch initiation. Furthermore, negative relationships were found between egg size and predation and duration of survival, respectively. We assume that concealed nests, early breeding and breeding experience diminish predation in salt marsh breeding redshanks. Thus, redshank reproduction appears to be affected by interactive effects of timing of breeding and vegetation facilitating early breeding. In contrast to open-nesting species, breeding in riparian habitats next to waterbodies may be disadvantageous for species breeding concealed in vegetation if these are covered by less structured vegetation.     

Intensity of livestock grazing in relation to habitat use by brown hares (Lepus europaeus)

There is very limited information concerning livestock (sheep and goats) and brown hare Lepus europaeus interaction when both coexist. The effect of the intensity of livestock grazing on seasonal habitat use by hares, in a typical Mediterranean rangeland, was evaluated using the pellet-count method. Lightly grazed pastures were less preferred by hares compared with moderately grazed ones, whereas ungrazed pastures were used less intensively than grazed ones. Because livestock grazing reduces the quantity of standing biomass proportionally to its grazing intensity, forage resource was not the driving force for pasture selection. The increased use of moderately grazed pastures by hares in relation to lightly and ungrazed ones, where vegetation was more abundant, could be attributed to their reduced herbage height and density. This behaviour is probably a tactic that hares follow for predator avoidance, because they are more likely to detect visually approaching predators when feeding in a biotope with a limited herbaceous layer. The conclusion of this research is that livestock and brown hare coexistence may be compatible and beneficial rather than competitive when stocking rates do not exceed grazing capacity, leading to the conclusion that proper livestock grazing and hare population management can be feasible in practice.     

Variation in structure and diversity in Mediterranean grasslands related to trophic status and grazing intensity

Abstract. The spatial organization of Mediterranean grassland in Spain is described, based on samples from 71 sites covering the existing variation in slope exposition and inclination.The whole set can be regarded as representing a trophic gradient, along which gradual variation in soil, species composition, biomass, and coverage were quantified. Corresponding to other studies from varying habitats, maximal species richness, diversity, and heterogeneity were observed on moderately infertile sites. Maximum species richness, over 60 species, occurred on sites with biomass values from 150–350 g / m². Species richness values are much higher and biomass values are much lower than those found in temperate grasslands. The decrease in diversity towards the mosteutrophic communities is stronger than expected, but can be easily explained by the high grazing pressure here. The variation in diversity observed runs parallel with that in heterogeneity. Zones with a high species richness also have a high heterogeneity, meaning a low amount of dominance. Grazing is understood as abiotic form of disturbance. Differences in grazing pressure may modify the relation between richness and fertility. While the greatest grazing pressure coincides with the most eutrophic communities, decreasing progressively towards the oligotrophic ones, the trend predicted by the resource availability theory is maintained; species diversity will be maximal at intermediate levels of disturbance. Absence of grazing in the eutrophic communities would lead to an investment in the soil of the unconsumed organic matter.     

Compatibility of livestock grazing with stand regeneration in Mediterranean holm oak parklands

Successful regeneration of holm oaks is the key to the conservation of the outstanding biodiversity levels in Spanish dehesa parklands. However, low densities of regeneration were measured in this study. The threshold for livestock stocking levels supporting regeneration was below all figures presently found in the dehesas. In the analysis of stand structure, a positive relationship between tree age and the age of agro-silvo-pastoral use of the dehesas was detected. This suggests that the forest cycle has been disrupted, and stands may dissolve gradually. Regeneration failure is an implicit component of this agroforestry system. An analysis of long-term abandoned dehesas situated at roadsides showed that holm oak stands are able to recover if grazing and cultivation are set aside. In a mail survey, managers of private large landholdings highly appreciated having holm oaks on their land, both for income- and non-income-related motivations, e.g. for the preservation of real estate value or family tradition. Land managers identified over-maturity of stands and regeneration failure among the top five problems of dehesas. Conservation policy should be directed towards incentive schemes, environmental education, and technical assistance.     

Effects of herbivorous birds on intertidal seagrass beds in the northern Wadden Sea

During autumn migration (September to December), brent geese (Branta b. bernicla) and wigeon (Anas penelope) feed on the seagrass Zostera noltii in the nearshore, upper tidal zone leeward of the island of Sylt (eastern North Sea). To graze on leaves and shoots above the sediment and on rhizomes and roots below, these birds reworked the entire upper 1 cm layer of sediment eight times within this 3-month period. In addition, brent geese excavated pits 3–10 cm deep by trampling in order to feed on below-ground phytomass. About 12% of the seagrass beds became pitted to an average depth of 4.5 cm. Using net exclosures, it was estimated that birds removed 34 g dry weight m^–2 of above-ground and 28 g of below-ground phytomass. This corresponds to 45% of the phytomass in September. Of the overall loss of phytomass from September to December, 63% was caused by birds. Roughly half of the leaves fell off anyway until December and the other half were taken by the birds. Below the ground, phytomass remained almost constant where birds were excluded, while with birds phytomass of rhizomes and roots was halved. In spite of this strong effect, in the next vegetation period the blade density was lower at former exclosure sites compared to the ambient seagrass bed. The underlying process seems to be a self-inhibition of dense overwintering seagrass by mud accretion. Assuming our experimental results can be scaled up to the entire seagrass bed, we hypothesize that in the sheltered upper intertidal zone, seasonal erosion caused by herbivorous geese and ducks is necessary for the persistence of Z. noltii. Received: 7 January 1999 / Received in revised form: 23 August 1999 / Accepted: 25 August 1999     

Effects of open grazing and livestock exclusion on floristic composition and diversity in natural ecosystem of Western Saudi Arabia

Livestock grazing is one of the main causes of rangeland degradation in Saudi Arabia. Fencing to exclude grazers is one of the main management practices used to restore vegetation and conserve biodiversity. The main objectives of this study were to investigate the changes in plant diversity and abundance, floristic composition and plant groups of the major life forms in response to thirty-five years of grazing exclosure in western Saudi Arabia. These vegetation attributes and palatability were compared in 30 sampling stands located in the excluded and grazed sites. Our results showed that livestock exclusion significantly increased covers, density and species richness of annuals, grasses, perennial forbs, shrubs and trees. Exclosure enhanced the abundance and richness of palatable species and depressed the development of weedy species. About 66.7% of the recorded species at the excluded site were highly palatable compared to 34.5% at the grazed site. In contrary, about 55.2% unpalatable species were found in the grazed site compared to 25.8% in the protected site. Jaccard’s similarity index between the excluded and grazed sites showed lower values of 0.39%, 0.40% and 0.31% at levels of families, genus and species, respectively. The results suggest that establishing livestock exclusion may be a useful sustainable management tool for vegetation restoration and conservation of plant diversity in degraded rangelands of arid regions.     

Reduced tuber banks of fennel pondweed due to summer grazing by waterfowl

In this study we investigated the effect of summer bird herbivory on the belowground tuber formation of fennel pondweed (Potamogeton pectinatus L.). Cumulative grazing pressure of four waterfowl species (mute swans, mallards, gadwalls and coots) in the summer was calculated based on timing of grazing and body mass of the grazers. The resulting grazing pressures were significantly negatively correlated with mean autumn tuber biomass in three of the four years of study. Moreover, summer grazing pressures explained more of the variance in tuber densities than water depth, sediment particle size distribution or any interactions of these variables did in the same three years. We propose that herbivory early in the summer has the most substantial impact on the clonal reproduction of macrophytes. Herbivores with a large body mass and early congregation for moulting may be the key waterfowl species in diminishing propagule biomass. Hence, they may present pre-emptive, time-staggered competition to consumers of the belowground biomass in autumn, such as migratory swans and diving ducks.     

Water use and feeding patterns of the marsupial western grey kangaroo (Macropus fuliginosus melanops) grazing at the edge of its range in arid Australia,as compared with the dominant local livestock,the Merino sheep (Ovis aries)

Western grey kangaroos (Macropus fuliginosus melanops) are large, foregut fermenting herbivores common to Australia's southern woodlands and shrublands, but they extend well into semiarid habitats at the north-eastern edge of their range. At this range boundary, western grey kangaroos occupy open chenopod (saltbush) shrubland, along with Australia's other large native kangaroos, as well as with extensive pastoral stock, primarily the wool-breed merino sheep. In this habitat, within a large naturally vegetated enclosure (16 ha), western grey kangaroos grazing sympatrically with merino sheep spent much of the day resting under shade trees, and fed mainly during the evening and early morning, mainly on grasses and flat-leaved chenopods. On this diet, western grey kangaroos had water turnovers similar to those of red kangaroos, at 1.1 L d⁻¹. Sheep, however, used 7.7 L of water each day. Thus, although the sheep were twice the average body mass of kangaroos, the sheep used more than seven times as much water. This level of water use by sheep was almost half that previously reported for sheep at the same site feeding mainly on salt-laden chenopods (ca. 12 L d⁻¹), but was consistent with other studies showing lower water usage by sheep feeding on trees and low-salt shrubs; foregut (rumen) contents of our sheep comprised 35% tree browse. Overall, our data do not support suggestions that western grey kangaroos are limited mainly by water at this arid range-boundary. Notably, the western grey kangaroos’ feeding behaviours were consistent with those of other arid-zone kangaroos, highlighting distinctive differences in the ecological physiology of the foregut fermenting kangaroos and the ruminant sheep.     

Germination and population structure of spear thistle Cirsium vulgare in relation to experimentally controlled sheep grazing

Summary The size and number of rosettes of Cirsium vulgare were censused in a 4 ha sheep grazing trial on lowland calcareous grassland in paddocks receiving controlled winter, spring and summer grazing treatments. Spring grazing significantly increased thistle rosette numbers, and there were always fewer rosettes in size classes 250mm–300 mm under the heavier grazing treatments. The emergence of seeds of Cirsium vulgare sown by hand into each of the grazing treatments was monitored and showed a positive effect of spring grazing. When the percent emergence of sown seeds was used as a covariate in the analysis of rosette numbers in the experimental paddocks, it was found to account for 77% of the variance in thistle numbers. It was concluded that the main effect of grazing on thistle rosette numbers is an indirect one exercised through the effects of grazing on germination conditions. Seeds of Cirsium vulgare were sown into artificial gaps at another grassland site, to determine the effect of gap-size upon seedling emergence. Germination was poor, but significantly more seedlings emerged in gaps 10–20 cm diameter, than in 5 cm gaps or in control plots without a gap. A computer simulation model was used to explore the relationship between gap density and thistle population dynamics. A threshold density of gaps was found to exist, below which thistle populations went extinct, and above which the thistle population grew geometrically. The degree of aggregation of dispersed seeds did not alter the threshold gap-density for plant extinction, but did affect the rate of increase of the thistle population when the threshold gap-density was exceeded. It is concluded that strategies for controlling Cirsium vulgare populations by grazing manangement will be most effective if aimed at reducing suitable sites for establishment in spring. The success of such attempts will depend upon the soil fertility of the site, and sudden outbreaks of Cirsium vulgare infestation can be expected if a sward gradually deteriorates through over-grazing.     

Variations in the mussel population of the Dutch Wadden Sea in relation to monitoring of other ecological parameters

The pattern of distribution of intertidal mussel beds is relatively constant over a number of years although their surface area can vary greatly. The abundance of mussels shows much greater fluctuations. In the western part of the Dutch Wadden Sea, west of the Terschelling tidal divide, the amount of mussels on natural beds fluctuated between 1 and 24 million kg fresh weight during the years 1949 to 1988. In the eastern part of the Dutch Wadden Sea the biomass varied between 5.5 and 180 million kg. The influence of the mussels on the ecosystem therefore can be very different between years. When many mussels are present the whole watermass can be filtered every few days. In years with few mussels present the filtering may take one month. It is argued that monitor programmes for a.o. nutrients, chlorophyll and growth rates of benthic organisms are of limited value if there is no indication about the total amount of mussels in the area. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

Herbivory resistance traits in populations of Poa ligularis subjected to historically different sheep grazing pressure in Patagonia

Seeding selected populations with high grazing resistance may foster recovery of plant populations threatened by overgrazing. Resistance to grazing depends on grazing avoidance (escape from grazers) and grazing tolerance (ability to growth after defoliation). Many studies of grazing tolerance defoliate plants at a fixed height instead of removing the same proportion of biomass and therefore confound tolerance with avoidance. For this reason, the information on evolution of tolerance to defoliation at the intraspecific level is remarkably scarce despite the abundance of papers published that evaluate responses to defoliation. The estimation of the cost of tolerance is also troublesome because current methods usually include spurious correlations due to correlation between variables that share common terms. The objectives of this paper were to assess the intraspecific variation in tolerance and in traits associated with avoidance and growth in populations with different sheep grazing histories. We also estimated the percentage of biomass removed when the defoliation treatment was imposed at fixed height in order to separate tolerance and avoidance. Finally, we estimated the cost of tolerance using a new method proposed for spurious correlations. Results of a greenhouse experiment indicated no difference in tolerance among the three compared populations. However, the populations from overgrazed fields had more prostrate growth form, higher specific leaf area, and higher tillering rate (when no defoliated) than populations from exclosures. We confirmed that fixed height defoliation would have removed a higher proportion of shoot biomass from taller than from shorter individual plants, confounding grazing tolerance and avoidance. Regarding the cost of tolerance, we found no differences from a null model of no cost, indicating that the evolution (or future breeding) of more tolerant genotypes would not be constrained by this cost.     

Plant community structure in relation to grazing and environmental changes along a north-south transect in the western Kalahari

Sixteen vegetation types were described from a north-south transect in the western Kalahari. Pronounced differences were found between communities on the nutrient poor red Kalahari sand, covering most of the area and those on fine soils and white calcareous sand. Syntaxa resulting from severe overgrazing by livestock were in most cases clearly distinguished from the less disturbed vegetation. The communities on red sand consisted mainly of shrub savanna dominated by perennial tufted grasses, whereas in the vegetation on calcareous material and on overgrazed land, forbs, dwarf shrubs and shrubs played a more important role. The major communities on red sand showed a clear geographical zonation roughly corresponding to the gradient in mean annual rainfall and its interannual variation. In the northern and central Kalahari these syntaxa were dominated by species of Sudano-Zambezian origin and in the southern Kalahari by species showing Karoo-Namib affinities.     

Relationships between graminoid growth form and levels of grazing by caribou (Rangifer tarandus) in Alaska

Herbivores and their forage interact in many ways, in some instances to the benefit or detriment of herbivore and vegetation. Studies of wildebeest (Connochaetes taurinus) in Africa and snow geese (Chen caerulescens) in the Arctic have suggested that these grazers enhance graminoid production in certain sites by repeatedly using them. Other studies have concluded that herbivores are sensitive to local variation in forage quality and quantity, and preferentially use those sites that are intrinsically more productive. In this study, caribou (Rangifer tarandus) were observed foraging at different densities on two adjacent Alaskan ranges, within which particular feeding sites contained predictably high, medium, or low densities of caribou. Vegetation from one high- and one low-use site on each of the high- and low-density ranges was sampled and monitored for productivity, measured as re-growth following clipping, with the objectives of determining which forage characteristics influence usage by grazers and whether the productivity and nature of graminoid growth after clipping were related to grazing history. Forage biomass density (g/m³), shoot density (number/m²), stand densities of nutrients and minerals (g/m³), and forage concentrations of nutrients and minerals (g/100 g tissue) correlated positively with use of sites by caribou. Productivity was independent of previous use by grazers, but consistent within ranges. These results indicate that caribou are sensitive to local variation in forage quality and quantity, preferentially use those sites with higher returns of nutrients and minerals, and have the potential to enhance graminoid growth on sites that are inherently more productive.     

Lack of intraspecific variation in resistance to defoliation in a grass that evolved under light grazing pressure

Grass species frequently show marked intraspecific variation inmorphology and tolerance to defoliation. Remarkably, most of this evidencecomesfrom grasslands with long evolutionary history of grazing. Here, we explore theintraspecific variation in grazing tolerance and morphometric traits associatedwith grazing avoidance of Paspalum dilatatum (Poir.), agrass from the Flooding Pampa (Argentina), where grazing is a novel disturbancein evolutionary time. We performed a clipping experiment in a greenhouse withtwo populations from sites with contrasting short term grazing regime:continuous grazing and 20 year-old grazing exclosure. The populations did notdiffer in their tolerance to clipping, and showed minor differences in the wayclipping affected plant height, a trait associated with grazing avoidance. Ourresults indicate that there are exceptions to the generalized findings of highlevels of intraspecific variation in grazing resistance among populations ofgrasses. These exceptions may be associated to evolutionary history of grazing.     

In situ grazing pressure and diel vertical migration of female Calanus euxinus in the Black Sea

Gut pigment and abundance of the female Calanus euxinus (Hulsemann) weremeasured from several water layers (defined by density values), with3–5 h intervals during 30 h and 21 h at a station in the southwesternBlack Sea in April and in September 1995, respectively. The female C.euxinus was observed to begin migration to the upper phytoplankton-richlayer approximately 3 or 4 hours before the sunset. Only a fraction of thefemale Calanus population (0.2% in April and 3.6% inSeptember) did not migrate but remained at the depth of the oxygen minimumzone during the nighttime. The migrating population was determined to havespent 7.5 h in the euphotic zone in April and 10.5 h in September. Thegrazing rate of female Calanus euxinus was measured from the gut contentdata collected from the layers which contain the euphotic zone. Thepercentage of primary production grazed by the female C. euxinus wascalculated as 14.5% in April and 9.5% in September. This revised version was published online in July 2006 with corrections to the Cover Date.